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The Enterobacteriaceae

The Enterobacteriaceae are a large, heterogeneous group of gram-negative rods whose natural habitat
is the intestinal tract of humans and animals. The family includes many genera (Escherichia, Shigella,
Salmonella, Enterobacter, Klebsiella, Serratia, Proteus, and others). Some enteric organisms, eg,
Escherichia coli, are part of the normal flora and incidentally cause disease, while others, the
salmonellae and shigellae, are regularly pathogenic for humans. The Enterobacteriaceae are facultative
anaerobes or aerobes, ferment a wide range of carbohydrates, possess a complex antigenic structure,
and produce a variety of toxins and other virulence factors.
The family Enterobacteriaceae have the following characteristics: They are gram-negative rods, either
motile with peritrichous flagella or nonmotile; they grow on peptone or meat extract media without
the addition of sodium chloride or other supplements; grow well on MacConkey's agar; grow aerobically
and anaerobically (are facultative anaerobes); ferment rather than oxidize glucose, often with gas
production; are catalase-positive, oxidase-negative, and reduce nitrate to nitrite; and have a 3959% G
+ C DNA content.
Morphology & Identification
Typical Organisms
The Enterobacteriaceae are short gram-negative rods. Typical morphology is seen in growth on solid
media in vitro, but morphology is highly variable in clinical specimens. Capsules are large and regular in
klebsiella, less so in enterobacter, and uncommon in the other species.
E coli and most of the other enteric bacteria form circular, convex, smooth colonies with distinct edges.
Enterobacter colonies are similar but somewhat more mucoid. Klebsiella colonies are large and very
mucoid and tend to coalesce with prolonged incubation. The salmonellae and shigellae produce
colonies similar to E coli but do not ferment lactose. Some strains of E coli produce hemolysis on blood
Growth Characteristics
Many complex media have been devised to help in identification of the enteric bacteria. One such
medium is triple sugar iron (TSI) agar, which is often used to help differentiate salmonellae and shigellae
from other enteric gram-negative rods in stool cultures. The medium contains 0.1% glucose, 1% sucrose,
1% lactose, ferrous sulfate (for detection of H
S production), tissue extracts (protein growth substrate),
and a pH indicator (phenol red). It is poured into a test tube to produce a slant with a deep butt and is
inoculated by stabbing bacterial growth into the butt. If only glucose is fermented, the slant and the butt
initially turn yellow from the small amount of acid produced; as the fermentation products are
subsequently oxidized to CO
and H
O and released from the slant and as oxidative decarboxylation of
proteins continues with formation of amines, the slant turns alkaline (red). If lactose or sucrose is
fermented, so much acid is produced that the slant and butt remain yellow (acid). Salmonellae and
shigellae typically yield an alkaline slant and an acid butt. Although proteus, providencia, and morganella
produce an alkaline slant and acid butt, they can be identified by their rapid formation of red color in
Christensen's urea medium. Organisms producing acid on the slant and acid and gas (bubbles) in the
butt are other enteric bacteria.
Salmonellae are motile rods that characteristically ferment glucose and mannose without producing
gas but do not ferment lactose or sucrose. Most salmonellae produce H2S. They are often pathogenic
for humans or animals when ingested. Arizona is included in the salmonella group.
The Salmonella-Arizona Group
Salmonellae are often pathogenic for humans or animals when acquired by the oral route. They are
transmitted from animals and animal products to humans, where they cause enteritis, systemic
infection, and enteric fever.
Morphology & Identification
Salmonellae vary in length. Most isolates are motile with peritrichous flagella. Salmonellae grow readily
on simple media, but they almost never ferment lactose or sucrose. They form acid and sometimes gas
from glucose and mannose. They usually produce H
S. They survive freezing in water for long periods.
Salmonellae are resistant to certain chemicals (eg, brilliant green, sodium tetrathionate, sodium
deoxycholate) that inhibit other enteric bacteria; such compounds are therefore useful for inclusion in
media to isolate salmonellae from feces.
Antigenic Structure
Enterobacteriaceae have a complex antigenic structure. They are classified by more than 150 different
heat-stable somatic O (lipopolysaccharide) antigens, more than 100 heat-labile K (capsular) antigens,
and more than 50 H (flagellar) antigens . In Salmonella typhi, the capsular antigens are called Vi
O antigens are the most external part of
the cell wall lipopolysaccharide and
consist of repeating units of
polysaccharide. Some O-specific
polysaccharides contain unique sugars. O
antigens are resistant to heat and alcohol
and usually are detected by bacterial
agglutination. Antibodies to O antigens
are predominantly IgM.
K/Vi antigens are external to O antigens
on some but not all Enterobacteriaceae.
Some are polysaccharides, including the K
antigens of E coli; others are proteins.
H antigens are located on flagella and are
denatured or removed by heat or alcohol.
They are preserved by treating motile
bacterial variants with formalin. Such H
antigens agglutinate with anti-H
antibodies, mainly IgG.
The classification of salmonellae is complex because the organisms are a continuum rather than a
defined species. The members of the genus Salmonella were originally classified on the basis of
epidemiology, host range, biochemical reactions, and structures of the O, H, and Vi (when present)
antigens. The names (eg, Salmonellatyphi, Salmonella typhimurium) were written as if they were genus
and species; this form of the nomenclature remains in widespread but incorrect use. DNA-DNA
hybridization studies have demonstrated that there are seven evolutionary groups. Nearly all of the
salmonella serotypes that infect humans are in DNA hybridization group I; there are rare human
infections with groups IIIa and IIIb. The species name Salmonella enterica has been widely accepted, and
the organisms in DNA hybridization group I are S enterica subspecies enterica. The organisms in the
other groups have other subspecies names. It seems probable that the widely accepted nomenclature
for classification will be as follows: S enterica subspecies enterica serotype Typhimurium, which can be
shortened to Salmonella Typhimurium with the genus name in italics and the serotype name in roman
type. National and international reference laboratories may use the antigenic formulas following the
subspecies name because they impart more precise information about the isolates .
There are more than 2500 serotypes of salmonellae, including more than 1400 in DNA hybridization
group I that can infect humans. Four serotypes of salmonellae that cause enteric fever can be identified
in the clinical laboratory by biochemical and serologic tests. These serotypes should be routinely
identified because of their clinical significance. They are as follows: Salmonella Paratyphi A (serogroup
A), Salmonella Paratyphi B (serogroup B), Salmonella Choleraesuis (serogroup C1), and Salmonella Typhi
(serogroup D). The more than 1400 other salmonellae that are isolated in clinical laboratories are
serogrouped by their O antigens as A, B, C
, C
, D, and E; some are nontypeable with this set of antisera.
The isolates are then sent to reference laboratories for definitive serologic identification. This allows
public health officials to monitor and assess the epidemiology of salmonella infections on a statewide
and nationwide basis.
Organisms may lose H antigens and become nonmotile. Loss of O antigen is associated with a change
from smooth to rough colony form. Vi antigen may be lost partially or completely. Antigens may be
acquired (or lost) in the process of transduction.
Pathogenesis & Clinical Findings
Salmonella Typhi, Salmonella Choleraesuis, and perhaps Salmonella Paratyphi A and Salmonella
Paratyphi B are primarily infective for humans, and infection with these organisms implies acquisition
from a human source. The vast majority of salmonellae, however, are chiefly pathogenic in animals that
constitute the reservoir for human infection: poultry, pigs, rodents, cattle, pets (from turtles to parrots),
and many others.
The organisms almost always enter via the oral route, usually with contaminated food or drink. The
mean infective dose to produce clinical or subclinical infection in humans is 10
salmonellae (but
perhaps as few as 10
Salmonella Typhi organisms). Among the host factors that contribute to resistance
to salmonella infection are gastric acidity, normal intestinal microbial flora, and local intestinal immunity
(see below).
Salmonellae produce three main types of disease in humans, but mixed forms are frequent
The "Enteric Fevers" (Typhoid Fever)
This syndrome is produced by only a few of the salmonellae, of which Salmonella Typhi (typhoid fever) is
the most important. The ingested salmonellae reach the small intestine, from which they enter the
lymphatics and then the bloodstream. They are carried by the blood to many organs, including the
intestine. The organisms multiply in intestinal lymphoid tissue and are excreted in stools.
After an incubation period of 1014 days, fever, malaise, headache, constipation, bradycardia, and
myalgia occur. The fever rises to a high plateau, and the spleen and liver become enlarged. Rose spots,
usually on the skin of the abdomen or chest, are seen briefly in rare cases. The white blood cell count is
normal or low. In the preantibiotic era, the chief complications of enteric fever were intestinal
hemorrhage and perforation, and the mortality rate was 1015%. Treatment with antibiotics has
reduced the mortality rate to less than 1%.
The principal lesions are hyperplasia and necrosis of lymphoid tissue (eg, Peyer's patches), hepatitis,
focal necrosis of the liver, and inflammation of the gallbladder, periosteum, lungs, and other organs.
Bacteremia with Focal Lesions
This is associated commonly with S choleraesuis but may be caused by any salmonella serotype.
Following oral infection, there is early invasion of the bloodstream (with possible focal lesions in lungs,
bones, meninges, etc), but intestinal manifestations are often absent. Blood cultures are positive.
This is the most common manifestation of salmonella infection. In the United States, Salmonella
Typhimurium and Salmonella Enteritidis are prominent, but enterocolitis can be caused by any of the
more than 1400 group I serotypes of salmonellae. Eight to 48 hours after ingestion of salmonellae, there
is nausea, headache, vomiting, and profuse diarrhea, with few leukocytes in the stools. Low-grade fever
is common, but the episode usually resolves in 23 days.
Inflammatory lesions of the small and large intestine are present. Bacteremia is rare (24%) except in
immunodeficient persons. Blood cultures are usually negative, but stool cultures are positive for
salmonellae and may remain positive for several weeks after clinical recovery.